the mission of the nevada agricultural experiment …naes.agnt.unr.edu/pms/pubs/50_2009_01.pdfthe...
TRANSCRIPT
The mission of the Nevada Agricultural Experiment Station is to develop
research capacity to support economic development in the state of Nevada,
especially in the agricultural sector, and to disseminate new research-
based information to stakeholders. In support of this mission, the Nevada
Agricultural Experiment Station offers outreach research publications for
use by agricultural producers throughout the state. This NAES Research
Bulletin provides peer-reviewed research findings that are applicable to
Nevada agriculture and based on excellent science. We trust you will find
this bulletin informative and useful.
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Wyoming Big Sagebrush
State & Transition Model
And Management Key for Nevada
First Approximation
Barry Perryman1, Sherman Swanson2, Gary Brackley3,
Brad Schultz4, Paul Blackburn5, Kent McAdoo6, Gary Medlyn7 and Robin Tausch8
INTRODUCTION
During the late twentieth century, public perception of Wyoming big
sagebrush (Artemisia tridentata ssp. wyomingensis) [Beetle & A. Young]
changed from being a ‘weed’ to a valuable resource in danger of extirpation in
some landscapes. The two polar perspectives, perceiving it to be of no value
and only competitive with grasses, or perceiving it to be so valuable and scarce
that we must never control it, neither serve or benefit land managers or the
wildlife that depend on this important habitat. During this period, a focus on
rangeland condition has shifted to a focus on ecological thresholds and the
information needed to allocate limited financial and other resources to those
1 Rangeland Ecologist, Nevada Agricultural Experiment Station and College of Agriculture, Biotechnology and Natural Resources Department of Animal Biotechnology, Reno NV 89557 2 State Range Specialist, University of Nevada Cooperative Extension and College of Agriculture, Biotechnology, and Natural Resources Department of Natural Resources and Environmental Science, 1000 Valley Rd. Reno NV 89512 3 State Range Conservationist, Natural Resources Conservation Service Nevada State Office, 1501 Longley Ln, Reno NV 89502 4 Humboldt County Educator, University of Nevada Cooperative Extension, 1085 Fairgrounds Rd., Winnemucca NV 89445 5 Soil Scientist, Natural Resources Conservation Service Elko Field Office, Elko, NV 89801 6 Northeast Nevada Natural Resources Specialist, University of Nevada Cooperative Extension, 1500 College Parkway, Elko, NV 89801 7 Soil Scientist, Bureau of Land Management Ely District, Ely NV 89315 8 Research Scientist, USDA Forest Service
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areas, times, and actions that are most important for maintaining rangeland
health.
This publication focuses on ecological sites capable of supporting plant
communities dominated by Wyoming big sagebrush. This shrub occurs at lower
elevations on valley bottoms, alluvial slopes, foothills, and mountain side slopes.
It typically inhabits areas too moist for salt desert shrub species and too dry for
mountain big sagebrush. While this subspecies is somewhat palatable to sheep
and mule deer, it is not as palatable as black sagebrush (Artemisia nova
[Nelson]) nor is it palatable to cattle.
This state and transition model and management key describes vegetation
change and management alternatives for the Natural Resources Conservation
Service ecological sites listed in Table 1. However, some areas where these
sites occur are better understood by disregarding their potential for transitioning
to a tree state because they are logistically far from those sites that generally
have juniper (Juniperus sp.) and/or pinyon pine (Pinus monophylla Torr. & Frem.)
trees. In general, the potential for transitioning to the tree state is greater for
Wyoming big sagebrush sites that have higher precipitation or elevation, have
deeper soils, or are closer to sites with these characteristics. Similarly, the
potential for transitioning to the tree state is greater for Wyoming big sagebrush
areas that have closer proximity to trees.
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Table 1. Ecological sites described by this model in Nevada (NRCS Ecological
Site Descriptions, 2003).
Site number Ecological site name Site number Ecological site name 023XY011NV Dunes 8-10" P.Z. 026XY099NV Coarse Loamy 8-10" P.Z. 023XY020NV Loamy 10-12" P.Z. 026XY100NV Stony Slope 10-12" P.Z. 023XY030NV South Slope 8-12" P.Z. 026XY102NV Gravelly Clay Loam 8-10" P.Z.023XY033NV Clayey 10-14" P.Z. 027XY007NV Loamy Slope 8-10" P.Z. 023XY038NV Droughty Loam 8-10" P.Z. 027XY008NV Droughty Loam 8-10" P.Z. 023XY039NV Loamy Slope 10-14" P.Z. 027XY029NV Gravelly Fan 8-10" P.Z.
Table 1 Continued Site number Ecological site name Site number Ecological site name
023XY040NV Granitic Fan 8-10" P.Z. 027XY045NV Sandy 8-10" P.Z. 023XY049NV Granitic South Slope 8-12" P.Z. 027XY051NV South Slope 8-10" P.Z. 023XY051NV Sandy 8-12" P.Z. 027XY054NV Loamy Slope 10-12" P.Z. 023XY057NV Granitic Loam 10-12" P.Z. 027XY058NV Loamy 10-12" P.Z. 023XY063NV Shallow Granitic Hill 10-14" P.Z. 027XY065NV Granitic Slope 8-10" P.Z. 023XY068NV Granitic Loam 8-10" P.Z. 027XY067NV Granitic Loam 8-10" P.Z. 023XY071NV Ashy Loam 10-12" P.Z. 027XY072NV Granitic Slope 10-12" P.Z. 023XY072NV Ashy Slope 10-12" P.Z. 027XY088NV Granitic Loam 10-12” P.Z. 023XY077NV Shallow Loam 10-14" P.Z. 027XY091NV Loamy Fan 10-12” P.Z. 023XY082NV Loamy Fan 10-12" P.Z. 027XY092NV Granitic Fan 10-12” P.Z. 023XY088NV Chalky Knoll 028AY005NV Sandy 8-10" P.Z. 023XY096NV Ashy Sandy Loam 10-12" P.Z. 028AY010NV Coarse Gravelly Loam 10-12" P.Z.023XY097NV Loamy Fan 8-10" P.Z. 028AY015NV Loamy 8-10" P.Z. 023XY099NV Channery Hill 8-10" P.Z. 028AY017NV Shallow Loam 8-10" P.Z. 023XY101NV Stony Slope 8-10" P.Z. 028AY022NV Gravelly Clay 8-10" P.Z. 023XY102NV Gravelly Clay Slope 10-12" P.Z. 028AY028NV Droughty Loam 8-10" P.Z. 024XY001NV Dunes 6-10" P.Z. 028AY031NV Loamy Fan 8-10" P.Z. 024XY005NV Loamy 8-10" P.Z. 028AY040NV Gravelly Loam 10-12" P.Z. 024XY006NV Dry Floodplain 028AY050NV Gravelly Clay 10-12" P.Z. 024XY013NV Loamy 10-12" P.Z. 028AY054NV Coarse Loamy Fan 8-10" P.Z.024XY017NV Sandy 8-10" P.Z. 028AY086NV Coarse Loamy Fan 10-12" P.Z.024XY020NV Droughty Loam 8-10" P.Z. 028AY091NV Loamy Fan 10-14" P.Z. 024XY026NV Stony Slope 6-10" P.Z. 028AY095NV Loamy 10-12" P.Z. 024XY028NV South Slope 8-12" P.Z. 028AY121NV Deep Loamy 8-10" P.Z. 024XY033NV Steep North Slope 10-12" P.Z. 028AY124NV Loamy Plain 024XY035NV Shallow Loam 10-14" P.Z. 028BY005NV Sandy 8-10" P.Z. 024XY045NV Eroded Slope 6-10" P.Z. 028BY007NV Loamy 10-12" P.Z. 024XY046NV Gravelly North Slope 028BY010NV Loamy 8-10" P.Z. 024XY047NV Shallow Loam 8-10" P.Z. 028BY014NV Loamy Plain 8-10" P.Z. 024XY058NV Sandy Loam 8-10" P.Z. 028BY045NV Loamy Fan 8-12" P.Z. 025XY013NV Churning Clay 8-12" P.Z. 028BY052NV Droughty Loam 8-10" P.Z. 025XY014NV Loamy 10-12" P.Z. 028BY054NV Silty Plain 8-10" P.Z. 025XY015NV South Slope 8-12" P.Z. 028BY056NV Silt Flat025XY019NV Loamy 8-10" P.Z. 028BY068NV Dune 8-10" P.Z. 025XY021NV Shallow Loam 8-12" P.Z. 028BY080NV Shallow Loam 8-10" P.Z. 025XY027NV Loamy 12-14" P.Z. 028BY082NV Loamy Fan 12+" P.Z. 025XY045NV Ashy Loam 8-10" P.Z. 028BY086NV Gravelly Clay 10-12" P.Z. 025XY066NV Ashy Loam 10-12" P.Z. 028BY094NV Calcareous Loam 10-14" P.Z.025XY070NV Loamy Fan 8-10" P.Z. 029XY006NV Loamy 8-10" P.Z. 026XY010NV Loamy 10-12" P.Z. 029XY010NV Loamy Slope 8-10" P.Z. 026XY011NV South Slope 8-10" P.Z. 029XY029NV Loamy 10-12" P.Z. 026XY015NV Shallow Loam 10-12" P.Z. 029XY049NV Sandy Loam 8-12" P.Z. 026XY016NV Loamy 8-10" P.Z. 029XY057NV Loamy Slope 12-14" P.Z. 026XY019NV Churning Clay 10-12" P.Z. 029XY073NV Bouldery Loam 8-12" P.Z. 026XY020NV Sandy 8-10" P.Z. 029XY075NV Loamy Slope 10-12" P.Z. 026XY022NV Stony Slope 8-10" P.Z. 029XY105NV Gravelly Clay 10-12" P.Z. 026XY024NV Droughty Loam 8-10" P.Z. 029XY106NV Gravelly Clay Slope 10-12" P.Z.026XY026NV Granitic Slope 10-12" P.Z. 029XY114NV Loamy Fan 8-10" P.Z. 026XY029NV Eroded Slope 8-12" P.Z. 029XY116NV Loamy Plain 026XY051NV Dune 8-10" P.Z. 029XY117NV Silty Plain026XY096NV Sandy Plain 029XY119NV Silt Flat026XY098NV Gravelly Loam 8-10" P.Z. 029XY158NV Coarse Loamy 8-10" P.Z.
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For the areas where this model and management key applies, we discuss two
management situations: 1) areas where cheatgrass (Bromus tectorum L.) and
other invasive weeds (annuals and perennials) are established and becoming, or
are already an important management factor; and 2) areas where natives are the
only ecologically important species established in the area or at least they still
dominate ecological processes and management concerns.
1) MANAGEMENT WITH CHEATGRASS AND OTHER INVASIVE WEEDS
On these landscapes, the presence of exotic and/or noxious, invasive weeds
threatens the natural resilience and utility of most if not all Wyoming sagebrush
plant communities. These species compete very effectively with native plants.
They can transition plant communities to new states (Fig.1) or dominate after
certain disturbances without appropriate and timely management action. Their
presence is always a hazardous situation. When common, their presence
typically results in a transition to a new state because the exotic weeds, not the
desired species, determine ecological processes.
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sorbitol positive
Introduction
PERENNIAL HERBACEOUS STATE
DIMINISHED PERENNIAL HERBACEOUS UNDERSTORY
PERENNIAL HERBACEOUS UNDERSTORY
MOSTLY ABSENT
REQUIRES ADDITIONAL INPUTS.
SHRUB STATE
PJ CONTROL AND NEW UNDERSTORY PROPAGULES.
DIMINISHED SHRUB S
TREE STATE
?
ALTERED SITE POTENTIAL STATE
WYOMING BIG SAGEBRUSH STATE & TRANSITION MODEL (WITH CHEATGRASS AND/OR OTHER INVASIVE WEEDS ON SITES CAPABLE OF SUPPORTING TREES)
SHRUB AND TREE
REMOVAL AND
SUCCESSFUL
SEEDING
SHRUB AND TREE
REMOVAL
ANNUAL GRASS FIRE CYCLE STATE
FIRE RESISTANT SHRUBS AND CHEATGRASS
CHEATGRASS AND OTHER ANNUALS
ACCELERATED
EROSION
FIRE OR PJ CONTROL WITHOUT
SUCCESSFUL SEEDING
?
PERENNIAL INVASIVE WEED STATE
EXPLOSION OF
PERENNIAL INVASIVE
SPECIES
SEEDED PERENNIAL HERBACEOUS STATE
FIRE OR FIRE SURROGATE
SUCCESSION
SEEDED HERBACEOUS WITH SAGEBRUSH
FIRE O
RP
jC
ON
T RO
L AN
D
S UC
CE S SF U
L SE E D
I NG
SUCCESSION
FREQUENT FIRESINTRODUCTION AND EXPLOSION OF
PERENNIAL INVASIVE SPECIES
REL
ATI
VE
AB
UN
DA
NC
E
EARLY SERAL DOMINANTS
HERBACEOUS PERENNIALS SAGEBRUSH OVERSTORY
DOMINANT TREE SPECIES
SAGEBRUSH UNDERSTORY
FIRE A
ND
WEED
EXPLO
SION
EXPLOSION OF
PERENNIAL
INVASIVE SPECIES
SUCCESSION
FIRE OR FIRE SURROGATE
SEEDED HERBACEOUS
NO
FIRE
UNDERSTORY MOSTLY ABSENTHERBACEOUS
PLANT COMMUNITY
SHRUBBY OVERSTORYWITH HERBACEOUS
UNDERSTORY
NO FIRE TOBALANCE GRAZING
ANNUAL GRASS FIRE CYCLE STATE AFTER FIRE AND WITHOUT SUCCESSFUL SEEDING
PERENNIAL INVASIVE WEEDS DOMINATE IF THEIR POPULATIONS ARE ALLOWED TO EXPLODE
NO FIRE
NO
FIRE
FACILITATED SUCCESSION
NO FIRE
Fig. 1. Each box is shown as a different color to identify that it is a different state. The solid-line arrows between boxes are irreversible transitions across thresholds. Dashed-line arrows show active restoration of ecological processes. Inset box shows relative abundance of plant groups and relative sequence of transitions through succession without proactive management.
Page 7 of 34
PERENNIAL HERBACEOUS STATE Description: The plant community is dominated by deep-rooted perennial
bunchgrasses, with perennial forbs and varying amounts of Wyoming big
sagebrush and other shrubs. Sagebrush can dominate the plant community and
juniper and/or pinyon pine trees may be present as seedlings, saplings, or sparse
mature trees as long as the understory remains robust. If the perennial
understory is dense and vigorous enough to recover quickly after being released
from the competition of woody plants, the vegetation has not crossed a threshold
to the shrub or tree state. Descriptions of the ecological sites listed in Table 1
provide relative species composition and production data for each ecological site
in this perennial herbaceous state. Cheatgrass (or other non-native annual
plants) is a minor component of the understory vegetation.
Successional trajectories: The perennial herbaceous state plant community is
resilient or cyclic because secondary succession processes and disturbance
regimes are functional. Periodic release of understory perennials from increasing
competition from sagebrush is facilitated primarily by fire. However, other causes
for widespread shrub die offs have been noted. Normal fire frequency is
approximately 50-100 years (Wright and Bailey 1982). Without woody plant
removal, the plant community transitions to the shrub state or if trees are present,
to the tree state. On drier sites juniper may increase and on more mesic sites,
pinyon may increase. As transition to shrubs or trees occurs, the proportion of
cheatgrass in the herbaceous understory increases while perennial herbaceous
species decline. Poor grazing management of large domestic and/or wild
herbivores can diminish the vigor and expression of palatable perennial
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herbaceous plants. Removal of deep-rooted species may leave only or primarily
Sandberg bluegrass (Poa secunda [J.S. Presl.]) or cheatgrass. This makes
summer moisture and other resources more available to non-palatable shrubs
and/or trees and accelerates and increases the likelihood of transition to the
shrub, tree, or annual grass fire cycle state.
Management strategies to maintain the state: Manage for vigor, density, and
diversity of perennial herbaceous species. Include sagebrush and other woody
species in management objectives as desired. However, manage for no more
shrub and young tree canopy cover than is appropriate for the site in order to
maintain a resilient understory. Management should not allow the plant
community to transition across a threshold to the shrub or tree state. Research
indicates that herbaceous biomass production begins to decline when sagebrush
cover reaches 5-7% and density begins to decline when sagebrush cover
reaches about 12-15%, depending upon site potential (Rasmussen and others
2001, Winward 1991). To increase the vigor and density of the native perennial
herbaceous plants, intervene with mechanical control measures, prescription
grazing, herbicides, or very judicious use of prescribed fire. Shrub and young
tree control should be practiced as woody plant cover increases toward threshold
levels. However, caution is advised because cheatgrass can erupt from a seed
bank soon after control of woody plants opens niches that a sparse understory
cannot rapidly fill. The winter-annual, cheatgrass, out competes perennial
seedlings in most years on all but the sandiest soils. To minimize die off of
perennial herbaceous plants and bare patches, woody plant management may
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be needed more frequently than where only native perennial plants occur in the
understory. Where soils are erodible, minimize soil surface disturbance.
Wherever treatments disturb soil, ensure that adapted perennial plants or seeds
are available to compete with cheatgrass given the specific treatment conditions,
such as seedbed preparation, grazing regime, etc.
Grazing management should be designed to foster perennial herbaceous
species in the community. Excessive, prolonged, or poorly timed grazing by
animals with herbaceous diet preferences can increase shrubs. Shrubs can be
decreased by relatively intense winter grazing by animals with shrub diet
preferences. Supplemental feeding to concentrate cattle for mechanical damage
controls sagebrush in small patches, especially when shrubs are dry or frozen
and brittle. To limit bare ground after future disturbances, grazing and other land
or vegetation management actions should not weaken the perennial herbaceous
community. Where perennial herbaceous understory is weak and shrub cover is
still well below maximum, consider using selective herbicides to manage
cheatgrass and/or adjusting grazing management to restore vigor and density of
desirable understory species several growing seasons prior to controlling shrubs.
Bare ground is more susceptible to accelerated erosion, and invasive plants
establish faster in open niches. Management to maintain the perennial
herbaceous state (prescribed grazing and periodic control of woody plants) is
much more cost effective than management to return to this state once a
threshold has been crossed (control of woody plants, weed control, reseeding,
and temporary rest from grazing).
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SHRUB STATE
Description: Shrub cover has increased and perennial herbaceous understory
cover has decreased across a threshold level. Deep-rooted, perennial
bunchgrasses are rare to absent in the understory. The cheatgrass component
varies from present to dominant in the herbaceous understory. This state is very
susceptible to invasion by annual weeds before and especially after fire or other
large scale disturbance. Wyoming big sagebrush and other shrubs dominate the
plant community. Juniper and/or pinyon pine trees may be established on the
site but do not yet dominate ecological processes.
Successional trajectories: Native herbaceous understory is diminished from
perennial herbaceous state levels and may be absent or nearly so when
sagebrush cover reaches its maximum for the site. The relative abundance of
cheatgrass in the understory increases as perennial grasses decline. Eventually
cheatgrass dominates the sparse understory but drives long-term community
change for both shrubs and herbaceous species. Because a threshold has been
crossed, removal of grazing pressure will not restore the native herbaceous
component. This will coincidently require fire or other shrub control measures.
However, burning or other woody plant control measures without reseeding will
not return a mix of deep-rooted bunchgrasses and other plants characteristic of
the perennial herbaceous state. Return to the perennial herbaceous state
requires shrub control, cheatgrass control, reseeding, and possibly additional
management, depending on site-specific conditions. Return to the perennial
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herbaceous state requires facilitated succession beginning with transition to the
seeded perennial herbaceous state.
Fire or other major disturbance will increase the abundance of cheatgrass
and other annuals. A decline of big sagebrush in the overstory, coupled with an
increase in cheatgrass density indicates a transitional pathway to the annual
grass fire cycle state. With increasing cheatgrass fuel loads, the threat of wildfire
increases due to better fuel continuity and much higher flammability of this fine-
stemmed, often evenly distributed, early-growing and early-drying annual grass.
This change in fuel characteristics indicates a transition to the annual grass fire
cycle state that is completed by an inevitable fire. If trees are present, fire is
delayed, and tree invasion is not controlled, the plant community will transition to
the tree state. At the landscape scale, the rate of transition largely depends on
the geographic extent of fires, and can be quite extensive (up to 100,000 ac or
more).
Management strategies: To maintain the shrub state, or at least sagebrush,
prevention of wildfire is critical. Strategies often include creation of green strips
or other fuel breaks to keep wild fires small so that all sagebrush habitats are not
lost at once. Prescribed grazing may be used to reduce fine and/or woody fuels.
To transition to the seeded perennial herbaceous state, apply shrub and
weed control in conjunction with reseeding operations. Shrub control measures
could include herbicide, mechanical, or shrub-consuming herbivore treatments,
the judicious application of prescribed fire or wildfire. After wildfire or other shrub
removal, reseeding becomes urgent. Seeding is absolutely required before or
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within the first fall or early winter after shrub control. Thereafter, competition from
a rapidly expanding cheatgrass population may prevent seeding success.
Reseeding requires cheatgrass control unless a very hot fire removed all but 0-3
cheatgrass seeds per square foot. It also requires appropriate seedbed
preparation, planting date, and follow-up management. Reseeding treatments
could include native perennials, grasses, forbs and shrubs, and/or adapted non-
native perennial species. Where soil stabilization following wildfire is a priority
objective, seeding non-native perennial grasses with high seedling vigor may be
the best option. Investigate the feasibility of facilitated succession, seeding
initially with adapted non-native grasses and later inter-seeding with adapted
native herbaceous and/or shrub species.
SEEDED PERENNIAL HERBACEOUS STATE
Description: The choice of species in the seed mix, species in the pre-existing
seed bank, and the growing conditions in the first few years after the seeding
largely determine the species composition of the seeding. On many Wyoming
big sagebrush sites, few species will do predictably well. Even for crested
wheatgrass (Agropyron cristatum L. Gaertner), the most commonly seeded and
most dependable species, moisture following seedings is sometimes insufficient,
especially on the driest sites with the greatest soil limitations. Because perennial
bunchgrasses provide a clumped fuel composed of coarser stems that stay
green longer than cheatgrass, they generally reduce fire spread rates, and the
fire interval is generally long enough to allow sagebrush to become well
established (unless seeding design or management precludes it). Although
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functionally quite similar to the perennial herbaceous state, the seeded perennial
herbaceous state is shown as a separate state because there is always an initial
loss of genetic diversity once seeding is required.
Successional trajectories: Seedings often begin with expression of early seral
species present in the seed bank such as annual forbs. As perennials and
shrubs become better established, they typically exclude or severely diminish all
but the most aggressive of these early seral species or limit them to small
disturbed areas. If sagebrush was initially established by seeding or from seeds
left in safe sites, it structurally dominates a seeding more quickly. If not,
recruitment occurs from the edges or from unburned or untreated shrub islands.
Other species colonization depends on the mechanisms of seed dispersal and
their success in finding favorable microsites, or on treatments designed to
facilitate succession through their establishment. Transition back to the
perennial herbaceous state involves replacing the seeded species with native
species and/or increasing species diversity including the addition of shrubs,
especially sagebrush.
Management strategies: Seeding size, shape, or amount of edge and
orientation with respect to prevailing winds, as well as fire management
strategies to leave shrub islands or create a mosaic, can influence the process of
sagebrush re-establishment. During and after seeding establishment, livestock
grazing can be used to encourage niches for sagebrush and other species that
may be present. Conversely, grazing can be discouraged or managed
conservatively to favor only those species more palatable to livestock. Once the
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seeding has been used to avoid the transition to an annual grass fire cycle state,
management and additional seeding can be used to facilitate succession toward
various species compositions. Natives can be interseeded but often do not
compete well with established species unless steps like soil disturbance are used
to open new niches. Often the focus for management is simply maintaining the
seeding. This requires maintenance of ecological processes. Appropriate
activities would include keeping the seeding from becoming so dominated by
shrubs or weeds that the seeded understory is unable to survive a fire, or
allowing sufficient grazing to maintain plant vigor and reduce wolf plants.
Management strategies described for the perennial herbaceous state also apply
to the seeded perennial herbaceous state. However, specifics of grazing
management may differ according to the needs of the seeded and other desired
species.
TREE STATE
Description: Juniper and/or pinyon pine has established on a site and has
caused a decline in understory (herbaceous and shrub) cover and production
due to extended fire return interval. Although trees generally establish under
shrub canopies, they can invade the perennial herbaceous, seeded perennial
herbaceous, and shrub states. Trees have assumed ecological dominance,
driving future ecological processes. Understory (herbaceous and/or sagebrush)
has decreased across a threshold level defined by its lack of resilience to a tree-
removing disturbance. Tree biomass now dominates the plant community, with
leaf and fuel biomass as much as seven to eleven times the levels of perennial
Page 15 of 34
herbaceous or shrub states. However, tree cover is highly concentrated, often
leaving large bare interspaces that are susceptible to rill erosion, especially on
drier sites. Cheatgrass is present and often dominates the understory as trees
mature. Although live cheatgrass density and vigor may be lower in the tree
state than in other states, its seed bank is often extensive.
Successional trajectories: Herbaceous and/or shrub understory is diminished
from previous state levels to almost absent where trees are mature and the site
fully stocked. Shrub leaf biomass declines to approximately 20-25% of potential
when tree leaf biomass approaches 50% of maximum potential for the site
(Tausch and West 1995). During this process a 10 % increase in tree cover can
result in a 50% decline in understory production. The degree of resilience of the
understory is determined in part by the tree-removing disturbance. A very hot
wildfire may remove remaining herbaceous species and their seed reserves
(indicating the threshold to the tree state has been crossed) while a more gentle
form of tree removal may release these species from the tree competition
(indicating that the threshold had not yet been crossed). Once one or more
thresholds have been crossed in getting to the tree state, return to the perennial
herbaceous state requires transition to the seeded perennial herbaceous state
first. This requires shrub and/or tree control, reseeding, and usually other
management actions such as weed control. Cheatgrass is present and its
proportion in the understory tends to increase as native understory species
decline. If the native perennial understory is absent or sparse, fire or other tree
control measures alone will not increase most herbaceous/shrub understory
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species. Rather, the cheatgrass seed bank will increase cheatgrass abundance
after the release from tree competition, and transition the site to the annual grass
fire cycle state. Major soil erosion events after fire from severe wind, or from
major precipitation events on moderate or steeper slopes, can trigger a transition
to an altered site potential state. Mature tree stands may increase this risk by
allowing rills to form in large bare interspaces.
Management strategies: To manage this state for continued tree production,
protection from wildfire is essential. However, as trees grow, fuel accumulates
and tree canopies grow closer to each other. This increases the likelihood of a
catastrophic fire spreading across the landscape. Thinning a stand reduces fuel
loads. However, larger bare interspaces increase erosion hazard. Continued net
fuel production on this type often increases the risk of fires in neighboring
woodland types including areas where trees are very old because fire was
historically rare or involved only single trees. Management plans designed to
break up the landscape scale continuity of fuels with firebreaks, greenstrips, or
imposed differences in vegetation structure serve to reduce the risk of large fires
that leave watersheds barren. Applying tree control and rehabilitation treatments
in smaller patches increases the likelihood of fires creating a diverse mosaic of
habitats. This reduces the cost of future fire fighting, increases the opportunity
for fire use, and increases sustainability of ecological processes.
To transition to the seeded perennial herbaceous state, apply tree and
weed control and seed adapted perennials. After successfully attaining the
seeded perennial herbaceous state, facilitated succession can return the site to
Page 17 of 34
the perennial herbaceous state. Tree control measures could include prescribed
fire, mechanical, prescribed grazing, or herbicide treatments. Restoration
requires the use of site-adapted grass, forb, and shrub species and methods. If
site stabilization is a priority objective, non-native perennial herbaceous species
may be the best option for revegetation. Rehabilitation is required in the fall or
early winter immediately following tree removal.
ANNUAL GRASS FIRE CYCLE STATE
Description: Cheatgrass and/or other annual grasses and forbs (e.g., mustards)
dominate the herbaceous community. Most perennial herbaceous species
cannot compete with the dense population of cheatgrass and are absent or
nearly so. Fire intervals often shorten to 2 to10 years. Sagebrush is generally
unable to survive and reproduce with this fire frequency. Sprouting, fire-tolerant
shrubs may form a shrub overstory where fires are too frequent for sagebrush
but infrequent enough to support sprouting shrubs.
Successional trajectories: This plant community is functionally an annual
grassland. Cheatgrass initially dominates the site following wildfire. Sprouting,
fire-tolerant shrubs are the only woody plants, and these shrubs may eventually
dominate structural aspects of the area if fires are not too frequent. Cheatgrass
and annual forbs become the dominant vegetation after frequent, repeated fires.
Poor grazing management can shift species composition toward less palatable
species decreasing vegetative cover, while increasing erosion hazard. The level
of risk has increased for the community to transition to a perennial invasive weed
state. If perennial invasive species such as knapweeds (Centaurea sp.) are
Page 18 of 34
introduced, the plant community could rapidly become dominated by these
species, crossing a threshold to the perennial invasive weed state. This makes
rehabilitation to a seeded perennial herbaceous state more difficult, even with
extensive and intensive inputs. Fires export nitrogen and frequent fires may shift
the plant community toward undesired species tolerant of low-nitrogen soils.
Repeated fires expose soil to erosion more often, facilitating transition to the
altered site potential state during severe hydrologic or wind events.
Management strategies: To manage this state for continual annual grass
production, apply proper grazing for annual grassland. Leave sufficient biomass
and time for seed production and soil protection while consuming sufficient fuel to
reduce fire risk. This can be challenging due to highly variable production and
timing of production among wet and dry years. In the wettest years, grazing may
consume abundant forage in only some pastures or use areas or, with sheep
herding, in fuel breaks. In the driest years forage may be essentially absent.
To transition to the perennial herbaceous state, cheatgrass control and
reseeding operations are required. Mechanical, chemical, and grazing
treatments can reduce cheatgrass seed production. If fire intolerant shrubs like
sagebrush are included in the seed mix, a fuel management strategy must be
employed to reduce fire danger to shrub seedlings. Prescription grazing and
green stripping can be used across a landscape to reduce fuel loads, fire size,
and frequency. Establishment of seeded perennial herbaceous species (such as
bunchgrasses) will also reduce fuel continuity, potentially reducing the rate of fire
spread and size.
Page 19 of 34
PERENNIAL INVASIVE WEED STATE
Description: Although this state has yet to be identified in Nevada, one or more
of the weeds listed on the Nevada noxious/invasive weed list, or a new invasive
weed, could dominate the herbaceous vegetation, competitively excluding native
perennial herbaceous dominants. Weeds may burn readily and typically exclude
sagebrush and/or pinyon and juniper trees. Their competitive advantage, in an
environment without diseases, or insects from their ancestral home allows them
to displace most other plants to form virtual monocultures. Initial weeds may
facilitate the establishment of even more competitive invasive weeds.
Successional trajectories: The risk of transitioning to the perennial invasive
weed state increases after transition to the Shrub, Tree, Annual Grassland Fire
Cycle, and the Altered Site Potential States. Risk increases as soon as invasive
perennial plants, such as one of several knapweed species, begin to colonize an
area unless they are eradicated immediately upon discovery. Otherwise, initial
colonization generally expands toward a monoculture. Experience in other parts
of the western United States demonstrates the highly competitive nature of some
invasive weeds. However, which species will be most competitive on each
ecological site, state, or phase is still unknown. As initial infestations change
species composition and/or soil characteristics and site potential, other weeds
will likely become more competitive, causing instability in species composition.
Many invasive weeds are competitive, but do not effectively protect soil from
erosion or they are highly flammable, leading to unprotected soils after fires,
increasing the risk of crossing a threshold to the altered site potential state.
Page 20 of 34
Management strategies: Invasive plant colonies should be eradicated
immediately upon discovery. Once invasive weeds dominate a site, the expense
of weed control, follow-up control, and revegetation treatments generally exceed
on-site economic returns. However, these management strategies are justified to
quarantine weeds in one area, reducing spread potentials. Herbicides,
prescribed grazing (goats), and/or hand grubbing should be used to eradicate
small populations. Where eradication is no longer possible, mechanical,
chemical, and/or biological controls such as insects or prescribed grazing should
be used to control or confine infestations. Weed control areas will require
reseeding with the most competitive of adapted (native or non-native) desired
species and careful post-seeding grazing management to reduce the risk and
consequences of reinvasion. They may also require periodic treatment for
residual weeds. For whole landscapes dominated by noxious weeds, there may
be little option other than biological control. Yet biological controls are not
available for many weed species.
To accomplish vegetation management objectives suggested for this state
and transition model, care should be taken to avoid facilitating the spread of
invasive weeds. Expansions to the road network and soil disturbances increase
bare areas where invasive weeds can more easily establish. Virtually every
invasive weed population is first a roadside weed before its population explodes.
Many weed infestations begin in areas disturbed by machines, and some of
these are for vegetation management purposes. To prevent weed infestations
from spreading, it is important to routinely scout for new invasive weeds,
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especially in areas likely to be initial colonization areas (roadsides, waters,
riparian areas, turnout areas, corrals, utility corridors, borrow pits, treatment sites
etc.). Managers may also, remove or alter stresses on native perennial species
that can aid expansion from an affected area.
ALTERED SITE POTENTIAL STATE
Description: Accelerated erosion has resulted in loss of topsoil, altered
hydrologic characteristics (i e. reduced infiltration and increased runoff), and
lowered water and nutrient storage capacity. These changes to the growing
environment have resulted in an altered ecological potential for the site. For
example, a Wyoming big sagebrush site may become a shadscale (Atriplex
confertifolia [Torr. & Frem.]), site. Lowered site potential means lowered
vegetation production, less soil protection, and increased soil loss until a new
equilibrium is reached.
Successional trajectories: The risk of transitioning to the altered site potential
state increases after transitioning to the shrub, tree, annual grass fire cycle or
perennial invasive weed states. The new site potential and the array of possible
plant species and successional trajectories greatly depend on the soil remaining
as the rate of soil erosion stabilizes. For very shallow soils, plants survive by
tolerating extended periods without available soil moisture or by sending roots
deep into rock fissures. Cheatgrass, a winter annual, survives drought as seeds
that do not germinate in some years and by developing seed early. On sites
where the topsoil has been eroded away, clayey subsoil often becomes exposed
at the surface. Roots must penetrate the heavy clay and tolerate any shrinking
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and swelling of the clayey soil during germination. Plants must then be able to
persist with less soil moisture than available within an intact, non-eroded soil.
Clayey sites are also susceptible to invasion by medusahead rye (Taeniatherum
caput-medusa L. Nevski.).
Management strategies: Because topsoil or even subsoil has been lost, return
of the native perennial herbs and shrubs characteristic of the perennial
herbaceous state depends on soil forming processes that are slow under most
conditions. The area should now be managed under the guidance provided by
the state and transition model and ecological site description for the new
ecological site if one is available that fits. It is expected that the species
composition and limited productivity of the vegetation established on the altered
site will have a low resiliency and minimal utility.
2) NATIVES ONLY
In these plant communities and landscapes, non-indigenous species are usually
not present. Only plant species native to the Great Basin are important in
ecological processes and management. If present, non-natives reflect a
disturbance of vegetation that has left an open niche that can be easily filled
through recovery of native vegetation. The general model (Figure 2.) may
describe historic ecological processes and is still relevant in some areas.
Page 23 of 34
sorbitol positive
Introduction
WYOMING BIG SAGEBRUSH STATE & TRANSITION MODEL
(NATIVES ONLY ON SITES SUPPORTING TREES)
SHRUB STATE TREE STATEPERENNIAL HERBACEOUS STATE
SUCCESSIONAL CONTINUUM
RESILIANT TWO-WAY MOVEMENT NON-RESILIANT,
FINANCIAL OR MANAGEMENT INVESTMENT REQUIRED TO MOVE VEGETATION BACK ACROSS A THRESHOLD
REL
ATI
VE
AB
UN
DA
NC
E
DININISHED HERBACEOUS UNDERSTORY
UNDERSTORY ABSENT
REQUIRESADDITIONAL INPUTS
REQUIRES ADDITIONAL INPUTS
DIMINISHED SHRUB
UNDERSTORY
?
EARLY SERAL DOMINANTS
HERBACEOUS
PERENNIALS
SAGEBRUSH (OVERSTORY)
DOMINANT TREE SPECIES (OVERSTORY)
ALTERED SITE POTENTIAL STATE
UNKNOWN SPECIES COMPOSITION
SUCCESSION
FIRE OR FIRE SURROGATE
NON-RESILIANT,
HERBACEOUS PLANT COMMUNITY
SHRUBBY OVERSTORYWITH HERBACEOUS
UNDERSTORY
UNDERSTORYABSENT
Figure 2. Each box is shown as a different color to identify that it is a different state. The solid-line arrows between boxes are irreversible transitions across thresholds. Dashed-line arrows show active restoration of ecological processes. Middle box shows relative abundance of plant groups and relative sequence of transitions through succession without proactive management.
Page 25 of 34
PERENNIAL HERBACEOUS STATE
Description: The plant community is dominated by deep-rooted perennial
bunchgrasses, perennial forbs, and varying amounts of Wyoming big sagebrush.
Sagebrush can dominate the plant community and juniper and/or pinyon pine
trees may be present as seedlings, saplings, or very sparse mature trees, as
long as the understory remains robust. If the perennial understory is dense and
vigorous enough to recover after being released from the competition of woody
plants, the vegetation has not crossed a threshold to the shrub or tree state.
Descriptions of the ecological sites listed in Table 1 provide relative species
composition and production data for each ecological site in this perennial
herbaceous state.
Successional trajectories: Plant community is resilient or cyclic because
secondary succession processes and disturbance regimes are functional. Life-
form dominance (species composition) is controlled primarily by fire, although
Aroga moth or other phenomena can also thin or kill patches of Wyoming big
sagebrush. Normal fire frequency is about 50-100 years (Wright and Bailey
1982). Without periodic woody plant removal, the plant community will transition
toward the shrub state or if trees are adjacent to the site, to the tree state. On
drier sites juniper may increase and on more mesic sites, pinyon may increase.
Following wildfire, sprouting shrubs may dominate but will be gradually replaced
by perennial bunchgrasses and sagebrush. If the area is devoid of big
sagebrush, it could be restored through time with seeds from surrounding areas
and could be re-established more quickly with seeding. Poor grazing
management of large domestic and/or wild herbivores can diminish the vigor and
Page 26 of 34
expression of deep-rooted perennial herbaceous plants leaving primarily
Sandberg bluegrass. This makes soil moisture and other site resources more
available to competitive shrubs and/or trees and accelerates and increases the
likelihood of the transition to the shrub or tree state. If the perennial understory is
too sparse or weak to recover quickly after being released by fire or other major
disturbance, the vegetation has crossed a threshold to the shrub or tree state.
Large fires that remove sagebrush leaving no islands and/or repeated fires that
remove succeeding generations before reaching reproductive age (about 5 yrs)
may create large landscape areas with few or no sagebrush plants for extended
periods.
Management strategies: To maintain the perennial herbaceous state, limit
development of shrub or tree cover to what is appropriate for a resilient
herbaceous understory on the site. Research indicates that herbaceous biomass
production begins to decline when sagebrush cover reaches 5-7% and density
begins to decline when sagebrush cover reaches about 12-15%, depending upon
site potential (Rasmussen and others 2001, Winward 1991). Intervene with
prescribed fire, herbicide, mechanical control measures, and/or prescription
grazing. Grazing can be managed to reduce stress to palatable species,
especially during the growing season, slowing the advance of woody species.
Grazing can also be used to accelerate the process of sagebrush recolonization
after a fire. Shrub decrease can be fostered by relatively intense winter grazing
by animals with shrub diet preferences. Concentration of livestock at feeding
sites can reduce shrub density through mechanical damage to sagebrush,
especially when shrubs are frozen or dry and brittle. Grazing prescriptions
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should strive to maintain the vigor of the herbaceous community. Where the
perennial herbaceous understory is weak and shrub cover is still well below
maximum, investigate the feasibility of reseeding or adjusting grazing
management to improve vigor and density of desirable species in the understory
several growing seasons prior to controlling shrubs. Management to maintain
the perennial herbaceous state is often much more cost effective than
management to return to this state once a threshold has been crossed.
SHRUB STATE
Description: Herbaceous understory cover has decreased below a threshold
level. Shrub cover has increased above a threshold level. Wyoming big
sagebrush and/or sprouting shrubs dominate the plant community. Perennial
understory vegetation, especially deep-rooted bunchgrass, is incapable of
recovery after fire.
Successional trajectories: Native herbaceous understory declines
substantially from perennial herbaceous state levels and trends toward absence
when sagebrush cover approaches its maximum. If trees are present and not
controlled, the plant community will transition to a tree state. Because a
threshold has been crossed, transition to the seeded perennial herbaceous state
requires fire or other shrub control measures, reseeding operations and follow-up
management. Removal of grazing pressure alone will not restore the native
herbaceous understory characteristic of the perennial herbaceous state or
reduce shrub abundance. Burning or other shrub or tree control measures alone
will not return the mix of deep-rooted bunchgrasses and other perennial
herbaceous plants because the seed bank and seed source have been depleted.
Page 28 of 34
Woody plant removal will release fire-adapted shrubs and create open areas for
early-seral species and increase erosion potentials.
Management strategies: To maintain sagebrush stands, prevent wildfire but
control junipers and/or pinyon pines as needed. To transition to the seeded
perennial herbaceous state, apply shrub control measures in conjunction with
reseeding. Shrub control measures could include prescribed fire, herbicide,
mechanical, or grazing. Because one or more thresholds have been crossed,
reseeding is essential after wildfire. Reseeding with appropriate seedbed
preparation, planting date, and other methods should include a mix of adapted
desired (native or non-native) grass, forb, and shrub species. Where perennial
herbaceous understory is missing and shrub cover is still well below maximum,
investigate the feasibility of reseeding several growing seasons prior to
controlling shrubs. The goal of re-establishing a desired herbaceous component
may require a multi-step approach through many years or decades.
TREE STATE
Description: Juniper and/or pinyon pine has established on the area due to
extended fire return interval. Although trees generally establish under shrub
canopies, they can invade both the perennial herbaceous and shrub states. The
understory (herbaceous and shrub) has decreased below a threshold level
because tree cover has increased above a threshold level. Trees dominate the
plant community, with leaf biomass and fuel buildup often to seven to eleven
times the level of the perennial herbaceous or shrub states. However, canopy
cover is concentrated, often leaving large bare interspaces with erosion rills. The
perennial understory can no longer respond to fire or other tree-removing
Page 29 of 34
disturbances because seed banks, seed sources, and antecedent plants have
been depleted.
Successional trajectories: The perennial herbaceous and/or shrub understory
declines from previous state levels to almost absent as trees attain mature height
at normal density. Shrub leaf biomass declines to about 20-25% of potential
when tree leaf biomass approaches 50% of maximum potential for the site
(Tausch and West 1995). Each 1% increase in tree cover can lead to about a
5% decline in understory production. Post fire vegetation is dominated by early
seral species and the limited number of species that had survived tree
dominance and fire or other tree removal. Because one or more thresholds have
been crossed, transition to the seeded perennial herbaceous state requires
reseeding and tree control. If native perennial understory is absent, fire or other
tree control measures alone will not increase most herbaceous/shrub understory
species to levels found in the perennial herbaceous state. Species from the
surrounding landscape with wind-blown seeds will be favored. Removing trees
will create open areas susceptible to invasive species, sagebrush, and fire-
tolerant shrubs and/or accelerated erosion. After fire, major soil erosion from
severe wind or major precipitation events on moderate or steeper slopes can
trigger a transition to an altered site potential state. Accelerated soil erosion can
also occur in large bare interspaces where rills can develop. This is more
common on arid sites.
Management strategies: To manage this state for continued tree production,
protection from wildfire is essential. However, as trees grow, fuel accumulates
and tree canopies grow closer to each other increasing the likelihood of a hot
Page 30 of 34
crown fire spreading across the landscape. Thinning to reduce crown cover, fuel
load, and fuel continuity is critical to long-term maintenance of a woodland plant
community. Continued net fuel production on this type often increases risk of fire
in neighboring woodland types, including areas where trees are much older
because fire was historically infrequent or lightening strikes caused only single-
tree fires. Management plans should be designed to break up the landscape
scale continuity of fuels with firebreaks, greenstrips, or imposed differences in
vegetation structure.
To transition to the seeded perennial herbaceous state, apply tree control
measures in conjunction with reseeding. Tree control measures could include
prescribed fire, herbicide, or mechanical treatments. Reseeding should include
adapted grass, forb, and shrub species and appropriate seedbed preparation,
planting date, and follow-up grazing management and weed control where
needed. Because one or more thresholds have been crossed, reseeding is
essential after wildfire.
ALTERED SITE POTENTIAL STATE
Description: Accelerated erosion has thinned or eliminated topsoil, altered
hydrologic characteristics, and lowered water and nutrient holding capacity.
These changes alter the ecological potential of the site. Thereafter, reduced
vegetation cover and infiltration rate increases erosion potentials reducing site
potential until a new equilibrium is established.
Successional trajectories: New site potential, species composition of the
revegetation seed mix, and subsequent successional trajectories greatly depend
on characteristics of the remaining soil. For very shallow soils, plants survive by
Page 31 of 34
tolerating extended periods without available soil moisture or by sending roots
deep into rock fissures. On sites where clayey subsoil becomes exposed at the
surface after topsoil has eroded away, plants must tolerate any shrink-swell
characteristics of a clayey soil. Roots must be able to penetrate a heavy soil and
they must be able to persist with less available soil moisture.
Management strategies: Because topsoil or even subsoil has been lost, return
of native perennial herbs and shrubs characteristic of the perennial herbaceous
state depends on soil forming processes that are very slow under most
conditions. The area should now be managed under the guidance provided by
the state and transition model and ecological site description most similar to the
altered site. It is expected that the species composition and limited productivity
of the vegetation established on the altered site will have low resilience and
minimal utility.
CONCLUSION
This state and transition model and management key is designed to help
managers recognize opportunities to influence Wyoming big sagebrush sites in a
positive manner. It can be used for thinking at the site specific or the landscape
scale. Management opportunities are identified by determining the state and
successional trajectory by examining the vegetation. Pathways toward
thresholds indicate a need for action to prevent a transition to an unwanted state.
Thus, the model and management key helps set short-term and long-term
management objectives. Usually these objectives call for restoring resilience to
the perennial herbaceous state, protecting woody states from fire, focusing weed
control on prevention of high impact invasive weed population explosions, and
Page 32 of 34
reseeding immediately after a shrub or tree state burns. Management actions
are less risky, less expensive, and more satisfying when and where important
biological diversity remains and before difficult species, dangerous fuels, or
accelerated soil erosion dominate ecological processes. That is, before crossing
a threshold.
Across a landscape, the model helps focus attention on the highest priority
areas where an important management action or change has become urgent.
Across most landscapes, there are hot spots where site specific management is
urgently needed. There are other areas where the vegetation will remain resilient
into the future and areas where the threshold has been crossed. Crossing the
threshold to another state, decreases resilience and increases vegetation
treatment costs.
In general, the risk of losing the perennial herbaceous state is the highest
priority. This state cycles among a variety of species compositions to which
many wildlife and other species have adapted, including the many sagebrush-
dependent species. In addition many other resource values are produced in one
or more of the seral phases of this state. Its natural resistance to transitioning
across a threshold due to its resilience following natural disturbances makes this
state a low-cost management objective. However, after the introduction of exotic
invasive weeds and a century of altered fire regimes, this state is often at risk
where it still remains. Its increasing scarcity, and the presence of invasive weeds
that can more easily dominate after transitioning to the shrub or tree state,
elevates its value and its priority for management. Where it no longer remains,
the seeded perennial herbaceous state is its closest alternative.
Page 33 of 34
However, management does not equal preservation without disturbance.
This state is maintained by periodic disturbance. Between fires, time, succession
and growing season herbivory on the understory favors shrubs and trees. Fires
are the way nature balances these stresses to favor the understory. The focus of
land management in the Wyoming sagebrush type is to use management tools to
simulate natural disturbances at the right times and with the right combination of
other actions.
Page 34 of 34
REFERENCES:
Bestelmeyer, Brandon T., Joel R. Brown, Kris M. Havstad, Robert Alexander,
George Chavez, and Jeffrey E. Herrick. 2003. Development and use of
state-and-transition models for rangelands. J. Range Management,
56(2):114-126.
Laycock, W. A. 1991. Stable states and thresholds of range condition on North
American rangelands. J. Range Management 44(5): 427-433.
Rasmussen, G.A., M.P. O’Neill, and L. Schmidt. 2001. Monitoring Rangelands:
Interpreting What You See. Utah State Cooperative Extension Pub. NR
503.40p.
Stringham, Tamzen K., William C. Krueger, and Patrick L. Shaver. 2003. State
and transition modeling: An ecological process approach. J. Range
Management, 56(2):106-113.
Tausch, Robin J. and Neil E. West. 1995. Plant species composition patterns
with differences in tree dominance on a southwestern Utah pinon-juniper
site. Pages 16-23 In: Shaw, Douglas W., Earl F. Aldon, and Carol LoSapio
tech. Coord. Desired Future Conditions for Pinon-Juniper Ecosystems;
Proc. Of the Symp. 1994 Aug. 8-12 Flagstaff, Az USDA FS Gen Tech
Rept. RM-258.
Winward, A.H. 1991. A renewed commitment to management of sagebrush
grasslands. P. 2-7. In: Management of the sagebrush steppe. Agri. Exp.
Stat. Spec. Rept. 880. Oregon State University, Corvallis, OR.